Article

New Middle Pleistocene dental remains from Qesem Cave (Israel)

Authors:
  • VIESID, Vienna School of Interdisciplinary Dentistry, Austria
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Abstract

Ongoing fieldwork at the Middle Pleistocene site of Qesem Cave has resulted in the discovery of several new hominin teeth. These include a right upper deciduous canine (dc1), a right lower first deciduous molar (dm1), a right upper third premolar (P3), a right lower second molar (M2), a left lower third molar (M3), and an incomplete tooth (represented only by a single root). The teeth come from different stratigraphic layers at the site and may cover a time span of up to 200 ka. These specimens represent different tooth classes than the previously reported teeth from the same site. The current study presents metric and morphological data on the new Qesem Cave teeth as well as a discussion of their taxonomic affinities. The deciduous teeth show some features which, tentatively, seem to depart from the general Neanderthal pattern. The P3 and M2 show relatively simplified occlusal morphologies and lack "mass-additive" traits. The Qesem Cave permanent teeth seem to largely conform to the recently defined Eurasian dental pattern. The relatively large M2 shows a clear, continuous midtrigonid crest, but lacks a hypoconlulid. The M3 shows numerous accessory crests and furrows on the crown surface and also shows a nearly continuous midtrigonid crest. Thus, like the previously reported teeth from Qesem Cave, the new dental remains show some features that seem more consistent with fossils of early H. sapiens from the sites of Qafzeh and Skhul and some features which appear to align them with the Neanderthals. Given the uncertainties regarding the phylogenetic polarity of several of these features, a conclusive taxonomic diagnosis remains elusive and must await the discovery of additional, more complete remains from the site.

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... By far the most important specimen from Italy is represented by the Ceprano partial cranium. The calvarium shows a series of primitive traits combined with a number of synapomorphies Freidline et al., 2012;Simmons et al., 1991;Vandermeersch, 1981Vandermeersch, , 1989 33. Qesem four deciduous teeth (lower di2, dm1, and dm2 and upper canine), nine permanent teeth (upper I2, C, P3, M3 and lower C, P3, P4, M2, M3), incomplete dental root long root of lower C; long root length, large occlusal polygon and a lack of Neanderthal-like features (P4); lack of a reduced distal lobe (di2); no buccolingual expansion (I2) anterior fovea continuous with the central basin (dm1); squared cervical outline, high and inwardly bent dentine horns, faintly expressed midtrigonid crest (dm2); continuous mid-trigonid crest, separating anterior fovea from central basin (lower M2); midtrigonid crest (discontinuous on occlusal enamel surface, but continuous in enamel-dentine junction) (lower M3); marked degree of labial convexity (I2) Fornai et al., 2016;Hershkovitz et al., 2011Hershkovitz et al., , 2016Weber et al., 2016 associated with the expansion of the brain in the Middle Pleistocene. In a detailed cladistic analysis by Mounier and Caparros (2015: Fig. 2) Ceprano clusters away from Sima de los Huesos and Neanderthals, but fails to form a unified sister clade with other Middle Pleistocene hominins. ...
... At Qesem, a total of four deciduous teeth (mandibular di2, dm1, and dm2 and maxillary canine), nine permanent teeth (maxillary I2, C, P3 and M3 and mandibular C, P3, P4, M2 and M3) and an incomplete tooth root were recovered from the stratigraphic sequence spanning 400-200ka (Falgu eres et al., 2016;Hershkovitz et al., 2016). With the exception of the maxillary I2 and M3 and the mandibular deciduous di2, the teeth are mostly around 300 ka or older. ...
... With the exception of the maxillary I2 and M3 and the mandibular deciduous di2, the teeth are mostly around 300 ka or older. A detailed description is provided by Hershkovitz et al. (2011Hershkovitz et al. ( , 2016, Fornai et al. (2016), andWeber et al. (2016). The mandibular dentition is characterized by small crown size and generally plesiomorphous traits. ...
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... The Qesem Cave (QC) Middle Pleistocene hominin site has yielded several deciduous and permanent teeth (Hershkovitz et al., 2011Hershkovitz et al., , 2015 Weber et al., 2015) associated to the Acheulo-Yabrudian Cultural Complex (AYCC) and dated to about 420e220 ka (Barkai et al., 2003; Gopher et al., 2010; Mercier et al., 2013; Falgu eres et al., 2015). A complete Homo sp. ...
... Further quantitative studies on other QC and Levantine teeth showed a complex picture for the dental remains, in terms of both size and morphology (Weber et al., 2015). In spite of previous works (Hershkovitz et al., 2011Hershkovitz et al., , 2015 Sarig et al., 2015; Weber et al., 2015), there is still much uncertainty about the taxonomic nature of the QC inhabitants. First, they are represented by isolated teeth only. ...
... Dm 2 -QC2 is a lower left deciduous second molar. It is part of the QC dental record and was found within Amudian deposits possibly older than 300 ka (Hershkovitz et al., 2011Hershkovitz et al., , 2015). A first description of this deciduous tooth, accompanied by a preliminary assessment of dm 2 -QC2 dimensions, was carried out by Hershkovitz et al. (2011) who stressed the mixed presence of similarities to the Late Pleistocene modern humans (i.e., Qafzeh/Skhul specimens) on the one hand and Neanderthal-like features on the other. ...
Article
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The Mid-Pleistocene Qesem Cave near Tel Aviv in Israel yielded several hominin teeth and abundant faunal and cultural remains. The geological sequences of the cave were dated to 420,000–200,000 years ago. In this contribution, we focus on the three lower postcanine teeth which are among the oldest material from the cave. We used both Geometric Morphometrics and qualitative observations on the outer enamel surface and the internal enamel–dentine junction to investigate shape and size variation in a sample of Early-to Late-Pleistocene fossils (Sangiran, Mauer, Bilzingsleben, Ehringsdorf, Qafzeh, Ohalo), Neanderthals, and geographically diverse recent humans. Our approach based on three dental traits from three tooth types is able to distinguish quite well between dental specimens from anatomically modern humans (AMH) and Neanderthals (NEA). It also confirms an intermediate morphology of Mid-Pleistocene specimens in general, and the close proximity of Ehringsdorf to NEA. While the Qesem premolars display an intermediate shape between NEA and AMH, their size is definitely modern-like. The Qesem molar features a morphology and size closer to NEA. A possible explanation is the evolutionary dissociation of size and shape in premolars, and molars that are morphologically closer to NEA than premolars. It can be noted that a Mid-Pleistocene hominin population was present in Southwestern Asia that shows some Neanderthal affinities, probably more than Mauer and Bilzingsleben, but less than Ehringsdorf. With the current data, however, we cannot confidently assign the Qesem teeth to any existing taxon, nor exclude that it is an autochthonous phenomenon in the Levant.
... The Qesem Cave (QC) Middle Pleistocene hominin site has yielded several deciduous and permanent teeth (Hershkovitz et al., 2011Hershkovitz et al., , 2015 Weber et al., 2015) associated to the Acheulo-Yabrudian Cultural Complex (AYCC) and dated to about 420e220 ka (Barkai et al., 2003; Gopher et al., 2010; Mercier et al., 2013; Falgu eres et al., 2015). A complete Homo sp. ...
... Further quantitative studies on other QC and Levantine teeth showed a complex picture for the dental remains, in terms of both size and morphology (Weber et al., 2015). In spite of previous works (Hershkovitz et al., 2011Hershkovitz et al., , 2015 Sarig et al., 2015; Weber et al., 2015), there is still much uncertainty about the taxonomic nature of the QC inhabitants. First, they are represented by isolated teeth only. ...
... Dm 2 -QC2 is a lower left deciduous second molar. It is part of the QC dental record and was found within Amudian deposits possibly older than 300 ka (Hershkovitz et al., 2011Hershkovitz et al., , 2015). A first description of this deciduous tooth, accompanied by a preliminary assessment of dm 2 -QC2 dimensions, was carried out by Hershkovitz et al. (2011) who stressed the mixed presence of similarities to the Late Pleistocene modern humans (i.e., Qafzeh/Skhul specimens) on the one hand and Neanderthal-like features on the other. ...
Article
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The Qesem Cave Middle Pleistocene hominin site has yielded a well preserved lower second deciduous molar (dm2-QC2), among several other human dental remains. It has been previously described by Hershkovitz et al. using traditional methods. In this study, we used multiple approaches in order to characterize the outer and inner morphology of dm2-QC2, namely a descriptive investigation of the inner morphology, analysis of the dental tissues, and comparative 3D geometric morphometric investigation of various aspects of the dental crown based on data gathered from μCT images. Dm2-QC2 was compared to a sample of 44 specimens, including recent and fossil modern humans, Neanderthals, and Homo erectus.
... In more recent studies, Qesem lower molars were demonstrated to be morphologically closer to Neanderthals , and the deciduous second molar QC2 clearly indicates the presence of Neanderthal features in the Chibanian of Southwest Asia . While it remains a possibility that Neanderthals were present in Qesem by at least 200 ka (and possibly as early as 400 ka; Hershkovitz et al., 2016), until more remains are available, the teeth from Velika Balanica represent the earliest record of Neanderthals in the Eastern Mediterranean. ...
... Velika Balanica indicates a co-occurrence in the Balkans of Neanderthal morphology and cultural transfer from Southwest Asia into the Balkans as early as 300 ka. The dates of Sima de los Huesos (~430 ka or MIS 12; Arnold et al., 2014;Demuro et al., 2019), Visogliano (500e350 ka;Falgu eres et al., 2008Falgu eres et al., , 2010, and Fontana Ranuccio (~450 ka; Biddittu et al., 1979;Segre and Ascenzi, 1984;Muttoni et al., 2009), all showing decidedly Neanderthal morphology (Martin on-Torres et al., 2012;Arsuaga et al., 2014Arsuaga et al., , 2015Zanolli et al., 2018), are older than Qesem (400e200 ka; Hershkovitz et al., 2016) and Velika Balanica (~300 ka) indicating that Neanderthals may have moved from the West of Europe towards the Balkans and the Near East. Quinaderived tool assemblages like the one found in Velika Balanica are not recorded at any earlier European sites and follow a geographic trail from the East to the West. ...
Article
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... Morphologically, Prado Vargas 1360 is similar to dm1 specimens from Qesem Cave, Nadale 1, Sima de las Palomas, and Roc de Marsal in the presence of four cusps, an anterior fovea on the occlusal surface and a tuberculum molare (Arnold et al., 2016;Bayle et al., 2009;Hershkovitz et al., 2016;Trinkaus and Walker, 2017). These discrete traits have been identified in some Neanderthal lower dm1 (Bailey, 2002;Bailey and Hublin, 2006). ...
... following the criterion outlined in footnote k andArnold et al. (2009). recent) inFaerman et al. (1994),Mizoguchi (2002),Arnaud et al. (2016),Hershkovitz et al. (2016), andTrinkaus and Walker (2017). ...
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... Morphologically, Prado Vargas 1360 is similar to dm1 specimens from Qesem Cave, Nadale 1, Sima de las Palomas, and Roc de Marsal in the presence of four cusps, an anterior fovea on the occlusal surface and a tuberculum molare (Arnold et al., 2016;Bayle et al., 2009;Hershkovitz et al., 2016;Trinkaus and Walker, 2017). These discrete traits have been identified in some Neanderthal lower dm1 (Bailey, 2002;Bailey and Hublin, 2006). ...
... following the criterion outlined in footnote k andArnold et al. (2009). recent) inFaerman et al. (1994),Mizoguchi (2002),Arnaud et al. (2016),Hershkovitz et al. (2016), andTrinkaus and Walker (2017). ...
... The various Levantine Late Middle/Late Pleistocene Homo groups and their likely intra-and interrelationships. This chart is based on the current and previous studies(1,(12)(13)(14)(15). The NR Homo group (red box) dominated the region throughout the Late Middle Pleistocene, migrated from the Levant to Eurasia to establish sister populations, and later interbred with H. sapiens that reached the Levant ~200,000 years ago. ...
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Marom and Rak claim, on the basis of a few mandibular features, that the Nesher Ramla (NR) Homo is a Neanderthal. Their comments lack substance and contribute little to the debate surrounding the evolution of Middle Pleistocene Homo. Limitations and preconceptions in their study prevented them from achieving resolution beyond a dichotomous interpretation of the NR as either a Neanderthal or a modern human.
... The systematic excavations at the site have yielded, to date, a significant set of hominin permanent and deciduous teeth. According to Hershkovitz et al. (2016), the Qesem hominids show no affinities with Homo erectus; rather, they are more similar to later Middle Palaeolithic populations of this region, including Neanderthal affinities. ...
Article
Full-text available
The behaviour and mobility of hominins are dependent on the availability of biotic and abiotic resources, which, in temperate ecosystems, are strongly related to seasonality. The objective of this study is to establish evidence of seasonality and duration of occupation(s) of specific archaeological contexts at late Lower Palaeolithic Qesem Cave based on the study of ungulate teeth. Combining individual ageing using dental eruption and replacement with variability measurement of tooth microwear, we estimated the seasonality of occupations at different levels of the site and their relative duration. Information about the diet of the ungulates and the habitats where they were hunted was also derived from tooth mesowear and microwear analyses. In the different tooth assemblages analysed, where the fallow deer was the most abundant herbivorous species, animals were selectively hunted in specific habitats. For example, the fallow deer individuals brought back to the Hearth area had a different diet than those found in other parts of the cave. The Hearth area seems to have been used seasonally, probably during short-term events and more than once a year. The other areas of the cave show different seasons of game procurement and different patterns of occupation of the site, possibly more than once a year or, alternatively, for a longer period. The data suggest that Qesem Cave was inhabited by human groups for a long period, perhaps not continuously, and their knowledge of the environment surrounding the cave allowed them to allocate specific game, most probably from distinct hunting grounds, to designated activity areas at the cave. We hypothesise that the difference in feeding locations might have been related to specific needs of the cave inhabitants (food, hide and marrow extraction), and thus, animals from different hunting grounds were wisely used to maximise the potential of specific habitats in the environment. This study is an additional testimony to the ingenuity of the cave inhabitants that allowed them to persistently use the cave for a prolonged period in a year.
... What triggered these vast, creative innovations at that time? A study of human dental remains from Qesem Cave (Hershkovitz et al., 2016) indicates that AYCC hominins were no longer part of the Homo erectus clade, but were rather more similar to later populations of the Levant and shared characters of both Homo sapiens and the Neanderthal. It has been suggested that the emergence of the AYCC occurred concomitant with the speciation of a new hominin lineage in the Levant some 400,000 years ago, and that both processes were triggered by the disappearance of elephants from this area (Ben-Dor et al., 2011). ...
Article
In this paper, I contend that children had a unique position in prehistoric social systems, functioning as primary assimilators of new technologies. Their role is especially crucial at significant turning points in history, due to a number of childhood-cognitive mechanisms that are activated in learning and playing while engaging in innovative activity. I suggest that these mechanisms developed as part of an evolutionary process that has enabled humans to better adapt to change and prosper. This line of thinking is demonstrated through a synthesis of evolutionary, cognitive-psychological models and a case study from the archaeological record of the Levantine late Lower Paleolithic. In this time, humans developed a set of creative innovations which had to be learned and assimilated, such as the innovative production of blades. I argue that these cultural changes were successfully assimilated by groups inhabiting the Levant due to the enhancement of well-established learning mechanisms, in which children played a significant role. This role might have given them a unique status in their group – as preserving old traditions practiced by their ancestors but also as active agents, part of a collective group effort of tackling present and future challenges.
... Overall, both premolars show rather simplified crown morphology and resemble recent H. sapiens in the absence of accessory tubercles and ridges, and the lack of cingulum and/or buccal grooves.The expression of discrete morphological traits is not particularly discriminative among Middle/Late Pleistocene Homo, but H. sapiens are generally characterized by simple occlusal morphologies and low frequencies (or lack) of mesial and distal accessory ridges, accessory marginal tubercles, transverse crests and buccal cingula and/or grooves(14,77,78). The Qesem P 3 also shows a relatively simplified occlusal morphology, lacking a continuous crest joining the buccal and lingual cusps(88), and less lingual cusp reduction than in Misliya-1. The occlusal surface of the North African Middle Pleistocene specimen from Rabat is complicated by the expression of a mesial accessory cusp (P 3 ) and a mesial accessory ridge (P 4 ). ...
Article
To date, the earliest modern human fossils found outside of Africa are dated to around 90,000 to 120,000 years ago at the Levantine sites of Skhul and Qafzeh. A maxilla and associated dentition recently discovered at Misliya Cave, Israel, was dated to 177,000 to 194,000 years ago, suggesting that members of the Homo sapiens clade left Africa earlier than previously thought. This finding changes our view on modern human dispersal and is consistent with recent genetic studies, which have posited the possibility of an earlier dispersal of Homo sapiens around 220,000 years ago. The Misliya maxilla is associated with full-fledged Levallois technology in the Levant, suggesting that the emergence of this technology is linked to the appearance of Homo sapiens in the region, as has been documented in Africa.
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This paper brings together methodological, theoretical, and empirical analysis into the framework of linguistic diversity. It reflects both historical and contemporary research by economists and other social scientists on the impact of language on economic outcomes and public policies. We examine whether and how language influences human thinking (including emotions) and behavior, and analyze the effects of linguistic distances on trade, migrations, financial markets, language learning, and its returns. The quantitative foundations of linguistic diversity, which rely on group identification, linguistic distances as well as fractionalization, polarization, and disenfranchisement indices are discussed in terms of their empirical challenges and uses. We conclude with an analysis of linguistic policies and examine the trade-offs between the development of labor markets and the social costs that they generate in various countries. (JEL N30, Z13)
... Thirteen human teeth have been discovered at QC and are described as closer to the later populations (e.g. Skhul/Qafzeh) of this region, rather than to Homo erectus, although they also bear some Neanderthal traits ( Fornai et al. 2016;Hershkovitz et al. 2011Hershkovitz et al. , 2016Weber et al. 2016). ...
... Thirteen human teeth have been discovered at QC and are described as closer to the later populations (e.g. Skhul/Qafzeh) of this region, rather than to Homo erectus, although they also bear some Neanderthal traits ( Fornai et al. 2016;Hershkovitz et al. 2011Hershkovitz et al. , 2016Weber et al. 2016). ...
Article
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This paper presents the results of a flint type analysis performed for the small assemblage of bifaces found at the Acheulo-Yabrudian site Qesem Cave (QC), Israel (420–200 kya), which includes 12 handaxes, three bifacial roughouts, one trihedral, and one bifacial spall. The analysed artefacts were measured and classified into flint types based on visual traits. Also, extensive fieldwork aimed at locating potential sources was carried out. The bifaces were then assigned to potential flint sources, using both macroscopic and petrographic data, and were compared with a large general sample (n = 21,102) from various typo-technological categories and from various QC assemblages, studied by the same analytic process. Our results show that while the site is located within rich flint-bearing limestone outcrops of the Bi’na Formation (Upper Cretaceous Turonian), which dominate the general sample, non-Turonian flint types dominate the biface assemblage. The presence of roughouts and complete handaxes, alongside the complete absence of bifacial knapping by-products, as well as the absence of a clear spatial distribution pattern of the bifaces throughout the site’s sequence, stresses the fragmentation of the bifacial chaîne opératoire and suggests that the bifaces were not produced at the site but, rather, were brought to the cave in their current state. The extremely low quantity of bifaces at QC, compared with the overall rich lithic assemblages, suggests that handaxes did not play a major functional role in the QC hominins’ everyday lives. It is therefore possible that the QC bifaces originated from older contexts, most likely Acheulian sites existing in the vicinity of the cave, as part of the habit of the QC hominins of collecting older, previously knapped artefacts.
... Qesem has also yielded 13 human teeth from different parts of the stratigraphic profile. Data provided by morphometrical analysis and three-dimensional (3D) scanning point to the fact that the teeth from Qesem are not of Homo erectus (sensu lato) but bear similarities with the Late Pleistocene local populations of Skhul and Qafzeh, as well as some Neanderthal affinities (50). Therefore, the human fossils may belong to a yet unknown local hominin lineage of the Levant. ...
Article
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Bone marrow and grease constitute an important source of nutrition and have attracted the attention of human groups since prehistoric times. Marrow consumption has been linked to immediate consumption following the procurement and removal of soft tissues. Here, we present the earliest evidence for storage and delayed consumption of bone marrow at Qesem Cave, Israel (~420 to 200 ka). By using experimental series controlling exposure time and environmental parameters, combined with chemical analyses, we evaluated bone marrow preservation. The combination of archaeological and experimental results allowed us to isolate specific marks linked to dry skin removal and determine a low rate of marrow fat degradation of up to 9 weeks of exposure. This is the earliest evidence of such previously unidentified behavior, and it offers insights into the socio-economy of the human groups who lived at Qesem and may mark a threshold to new modes of Palaeolithic human adaptation.
... Finally, the systematic excavations at the site have yielded, to date, a significant set of hominin permanent and deciduous teeth. According to Hershkovitz et al. (2016), the Qesem hominids show no affinities with Homo erectus; rather, they are more similar to later Middle Paleolithic populations of this region (e.g., Skhul/ Qafzeh), although Neanderthal affinities also are present. ...
Article
The presence of fast-moving small game in the Paleolithic archaeological faunal record has long been considered a key variable to assess fundamental aspects of human behavior and subsistence. Birds occupy a prominent place in this debate not only due to their small size and to the difficulties in capturing them (essentially due to their ability to fly and their elusiveness), but also due to their possible role in the symbolic array in regard to non-nutritional elements (feathers, talons, etc.) and as reflectors of complex humaneworld relationships. In this study, we attempt to contribute to this topic by presenting taphonomical data of bird specimens from Qesem Cave (Israel), dated between 420 and 200 ka. Human-induced damage, including cut marks, peeling and human gnawing, has been identified on wing bones of Cygnus sp., Columba sp., Corvus ruficollis and Sturnus sp. Our evidence suggests that avian exploitation was not limited to food onlydeither to complement the human diet or as occasional food itemdbut also presumably for the use of feathers. While the consumption of birds as a dietary source seems to be evident as early as the Early Pleistocene, the non-alimentary use of inedible elements, such as feathers and talons, appears to be a practice from the Middle Paleolithic onwards. We argue that the combined nutritional and symbolic use of birds is one characteristic of the new mode of adaptation practiced already by the late Lower Paleolithic Acheulo-Yabrudian hominins in the Levant starting 400 ka. The Qesem findings point to the possible emergence of new cognitive and behavioral skills, which are followed in later periods in the Old World. Finally, we discuss the possible ontological and cosmological significance of humanebird interactions to illuminate our hypothesis regarding the emergence of a new perception of human relationships with the world as an integral part of the new Acheulo-Yabrudian mode of adaptation.
... Note the grooves separating the accessory cusp from the primary cusps. cation (Bailey, 2002;Bailey and Hublin, 2006;Glatz et al., 2008;Benazzi et al., 2011;Hershkovitz et al., 2016). Some argue that more complex occlusal morphology of the maxillary premolars, including accessory ridges and cusps, is characteristic of Neanderthals compared to anatomically modern humans (Benazzi et al., 2011). ...
Article
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The present study assesses the global distribution of marginal accessory cusps of the maxillary premolars. This trait, despite constituting one of the variables standardized by Turner and col-leagues (1991), has received little attention in morphological studies. Frequencies were calculated from data sheets collected by Christy G. Turner II for mesial, distal, and mesial + distal grades. Different geographic patterns were identified for both types of expression on the upper premolars. The patterned geographic distribution of these traits indicates their utility in biodistance investigations. In addition, the distinction between mesial and distal accessory cusps specified by Scott and Irish (2017) is recommended, as these two traits exhibit different geographic patterns.
... New discoveries of and research on human fossils in Africa and surrounding regions, have pushed back the timing of the origin of modern humans to a point earlier than that suggested by genetic analysis on one hand, and unveiled more evidence of the interbreeding of early modern humans and other hominid groups on the other. Thirteen 400-200 ka-old teeth found in Qesem Cave, Israel exhibit morphological features similar to the early modern human fossils discovered at Qafzeh and Skhul instead of Neanderthals (Hershkovitz et al., 2011(Hershkovitz et al., , 2016. Furthermore, hominid fossils discovered recently at Jebel Irhoud in Morocco have expanded the emergence of early modern humans in this region to around 300 ka. ...
Article
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Hypotheses concerning the origins of modern humans have been intensively debated, and two competing models, the recent “Out-of-Africa” and “Multiregional Evolution” paradigms, have dominated research and discussions for decades. Evidence from China has played a fundamental role in this debate: regional continuity and replacement by populations in-migrated from Africa have both been suggested and supported mainly by paleoanthropologists and geneticists, respectively. As more evidence has accumulated, new results obtained, and more scholars from various disciplines become involved, supporters of the recent “Out-of-Africa” model agree more or less with the “Multiregional Evolution” model regarding the complex history of modern humans and their interbreeding with other archaic populations (e.g., Neandertals). Recent discoveries of new human fossils, Paleolithic archaeological materials, and ancient DNA evidence in China have yielded a large body of information regarding the formation and development of modern humans in this region. However, controversies continue, including that most molecular biologists insist on the replacement of archaic populations by modern humans dispersed from Africa, while most paleoanthropologists and archaeologists propose an enhanced “Continuity with Hybridization” model. In this paper, we compile new results and progress in China and present the current debates and issues on the origins of modern humans. Finally, we offer several suggestions for future studies.
... Još uvek nije jasno ko je bio nosilac ovih promena. Jabrudijenska i ašelojabrudijenska nalazišta vezuju se za još uvek neidentifikovane hominine (bliske modernim ljudima iz Skula i Kafzeha) koji su potvrđeni u Kesemu, datovanom u period pre skoro 400 hiljada godina (Hershkovitz et al. 2011); lobanja iz Zutijeha, koja se vezuje za jabrudijen i koja je okvirno datovana u vreme pre 200-280 hiljada godina (MIS 7-8), takođe ne poseduje neandertalske crte (Hublin 2002), dok se šarantijenske industrije u Krapini javljaju sa brojnim ostacima neanderta-laca. Stoga se može pretpostaviti da je u jednom momentu, na nekom mestu došlo do transfera tehnologije između različitih vrsta hominina. ...
... The basic morphometric study indicates that the Qesem teeth are clearly not of Homo erectus (sensu lato). It has highlighted that while some of the traits are more Neanderthal-like, they are generally similar to the Late Pleistocene local populations of Skhul and Qafzeh caves dated to ca. 100 ka, (Hershkovitz et al. 2011Hershkovitz et al. , 2016). A 3-D scan of some of the teeth and various subsequent analyses resulted in similar conclusions, although Neanderthal affinities were more emphasized in some of these teeth (Fornai et al. 2016;Weber et al. 2016). ...
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Qesem Cave is a Middle Pleistocene site in Israel occupied between 420 and 200 ka. Excavations have revealed a wealth of innovative behaviors most likely practiced by a new hominin lineage. These include early evidence for the habitual and continuous use of fire, the repeated use of a central hearth, systematic flint and bone recycling, early blade production technologies, social hunting strategies and meat-sharing practices, and more. Fire was used throughout the 200,000 years of human occupation of the cave primarily for meat roasting and cooking. Roasting and cooking, we argue, had an important role in providing the necessary caloric intake of the cave’s inhabitants. We see fire as an essential element of the new post-Acheulian human adaptation in the Levant. The ample recurring evidence for focused and repeated use of fire for dietary purposes suggests that fire production, control, use, and maintenance were habitually practiced by the cave’s inhabitants and that fire-induced calories became central for their survival. We present an integrative view regarding the use of fire at Qesem Cave and discuss the role of fire within the framework of the significant cultural and biological transformations that took shape in the post-Acheulian Levant during the Middle Pleistocene.
... In ATD6-112, the groove pattern is less conspicuous due to the dental wear, but it is possible to observe a contact between the metaconid and the hypoconid. Although the posterior part of the crown is damaged, the morphology in this area seems to be similar to that of show a similar area ( Grine, 1984;Hershkovitz et al., 2016;Leakey & Wood, 1973, 1974Toro-Moyano et al., 2013). The tuberculum molare is particularly conspicuous in the TD6 dm 1 s. ...
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Objectives: During the last 13?years, the late Early Pleistocene Gran Dolina-TD6-2 level (Sierra de Atapuerca, northern Spain) has yielded an additional sample of 26 dental specimens attributed to Homo antecessor. In this report, we present a descriptive and comparative study of the six deciduous teeth. Methods: We provide external and internal morphological descriptions following classical terminology, as well as the mesiodistal and buccolingual measurements of the teeth. The internal morphology was described by means of micro-CT technique. Results: The TD6 deciduous teeth preserve primitive features regarding the Homo clade, such as the presence of styles in lower and upper canines and developed anterior and posterior foveae in the dm2 . However, other features related to the complexity of the crown morphology (e.g., cingulum) are not present in this sample. Furthermore, the great reduction of the talonid of the dm1 s is also noteworthy. Despite the limited comparative evidence, the presence of a remarkably well-developed tuberculum molare in the dm(1) and dm1 s from TD6 can be also considered a derived feature in the genus Homo. The TD6 hominins exhibit dental dimensions similar to those of other Pleistocene hominins. The dm1 s are buccolingually elongated and the buccolingual diameter of ATD6-93 is the largest recorded so far in the Homo fossil record. Conclusions: This study expands the list of plesiomorphic features of H. antecessor, and provides some information on the evolutionary status of this species. However, the identification of some advanced traits evinces a step towards the derived morphology of European Pleistocene teeth. The study of the deciduous dentition confirms the mosaic pattern of H. antecessor morphology revealed in previous studies of this hominin sample.
... The Middle Pleistocene site of Qesem Cave in Israel is dated between 420 and 200 kya and is culturally associated with the late Lower Palaeolithic Acheulo-Yabrudian Cultural Complex (AYCC). Ongoing research at the cave provides a wealth of well-preserved evidence for innovative behaviours throughout the sequence, including stone tools technology and use [19][20][21] , subsistence economy 22,23 , and site organisation 24,25 , possibly practised by a new hominin lineage 26 . ...
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For a long while, the controversy surrounding several bone tools coming from pre-Upper Palaeolithic contexts favoured the view of Homo sapiens as the only species of the genus Homo capable of modifying animal bones into specialised tools. However, evidence such as South African Early Stone Age modified bones, European Lower Palaeolithic flaked bone tools, along with Middle and Late Pleistocene bone retouchers, led to a re-evaluation of the conception of Homo sapiens as the exclusive manufacturer of specialised bone tools. The evidence presented herein include use wear and bone residues identified on two flint scrapers as well as a sawing mark on a fallow deer tibia, not associated with butchering activities. Dated to more than 300 kya, the evidence here presented is among the earliest related to tool-assisted bone working intended for non-dietary purposes, and contributes to the debate over the recognition of bone working as a much older behaviour than previously thought. The results of this study come from the application of a combined methodological approach, comprising use wear analysis, residue analysis, and taphonomy. This approach allowed for the retrieval of both direct and indirect evidence of tool-assisted bone working, at the Lower Palaeolithic site of Qesem Cave (Israel).
... Neandertals' deciduous teeth are less studied than permanent teeth for which numerous useful protocol have been developed (Bailey, 2004;G omez-Robles et al., 2008;Benazzi et al., 2011a). Though permanent teeth have received more attention than deciduous teeth in the last years several contributions have started to fill the gap (Grine, 2005;Bailey and Hublin, 2006;Macchiarelli et al., 2006;Bayle, 2008;Bayle et al., 2009aBayle et al., , 2009bBayle et al., , 2010Mahoney, 2010Mahoney, , 2013Toussaint et al., 2010;Zanolli et al., 2010;Benazzi et al., 2012Benazzi et al., , 2014aBenazzi et al., , 2015Macchiarelli, 2013;Fornai et al., 2014Fornai et al., , 2016Bailey et al., 2014aBailey et al., , 2014bBailey et al., , 2016Hershkovitz et al., 2015;Weber et al., 2016). Nonetheless, further work is required to understand the morphological and morphometric variability of hominin deciduous teeth. ...
Article
Despite new discoveries of human fossil remains, some aspects of paleoanthropological research are biased by the poor sample size, which limits our understanding of intra-species variability among the different hominin species. In this context, continuous assessment and reassessment of human fossil remains discovered decades ago, and often unknown to the scientific community, represent an opportunity to address this issue. Moreover, deciduous teeth are less studied than permanent dentitions, an aspect which contributes to limit our understanding. In the present study, we provide a detailed description of Tagliente 3 (upper right second deciduous molar) and Tagliente 4 (lower left deciduous canine), two deciduous teeth from Riparo Tagliente (Stallavena di Grezzana, Verona) attributed to Homo neanderthalensis. In terms of morphology and size, Tagliente 3 presents typical Neandertal derived features (e.g., likely large hypocone and complex topography of the enamel-dentine junction). Although deciduous canines usually do not provide substantial morphologically diagnostic information, Tagliente 4 falls in the upper range of the Neandertal variability for its bucco-lingual diameter. In terms of tissue proportions both teeth fall within the Neandertal range of variation: Tagliente 3 for the enamel thickness distribution and Tagliente 4 for the volume of the crown dentine. This work contributes to increase our knowledge on the variability of Neandertal deciduous dentition.
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To present a new dental specimen that will provide additional evidence for a better understanding of early European Upper Pleistocene hominin morphological variability. We described the morphology of this human right lower third molar at both the outer enamel surface and the enamel–dentine junction by means of micro‐computed tomography. In order to better understand hominin diversity, our morphological and metrical results were compared with those of other hominins obtained from published research. We provide a direct aspartic acid racemization dating of the molar. The direct dating (104.3 ka) situates the molar within the Marine isotopic stage 5d. The crown dimensions are comparable to those of the Sima de los Huesos sample and modern humans. The combination of a continuous middle trigonid crest and a well‐developed anterior fovea lies within the range of morphological variation reported for Neanderthal lower molars. The distal portion of the molar has a prominent protostylid. Crown and root morphology of this molar fits within the Neanderthal morphological pattern. However, both its dimensions and the absence of a hypoconulid tend to position this specimen away from contemporaneous Neanderthals and rather relate it more closely to some Middle Pleistocene populations. A new dental specimen is added to the Iberian Peninsula fossil record from the Marine isotopic stage 5, attesting to some degree of dental variability in the early Upper Pleistocene. Different views of the crown of CA‐2019‐E2‐2: (a) Photographs in distal (D), lingual (L), mesial (M), buccal (B) and cervical (C) views. (b) Photograph in occlusal view (left) and 3D surface model in occlusal (middle) and oblique (right) views.
Chapter
In this chapter we explore the period of modern human emergence between 300 and 100 ka. It was during this interval that some hominin populations began to develop “modern” human anatomical features and cultural behaviors. We review specific characteristics that distinguish Homo sapiens from previous forms and examine the physical evidence as to where and when these anatomical traits first appeared. With modern human anatomy came an array of technological innovations and behavioral changes. Elaborate core reduction strategies were invented, providing a window into our species’ developing cognitive functions. Different forms of symbolic expression and ritual activities suggest that humans had entered a more complex evolutionary stage of thought and perception. By the end of the chapter, we will have assembled enough clues to begin to address one of the fundamental questions of this book: what makes us human?
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Recycling is defined as a process in which waste materials can again become usable. In the common belief of many peoples, recycling is only considered a contemporary manifestation linked to the economic and ecological politics of industrialized societies. Both archaeological and historical records, however, prove that recycling has its roots back in time, being a common behavior of our ancestors as well as of many past societies. At the Late Lower Paleolithic site of Qesem Cave, Israel, research has identified a particular lithic trajectory oriented towards the production of small flakes by means of recycling, in the exploiting of old discarded flakes to be re-used as cores. The high density of this specific production throughout the stratigraphic sequence of the cave demonstrates that lithic recycling was a conscious and planned technological choice aimed at providing small and sharp items, most probably in order to meet specific functional behaviors. This particular lithic behavior persisted for some 200 kyr of human use of the cave and is not related to any shortage of flint, as the vicinity of the cave is exceptionally rich in flint sources. The exceptional conservation of use-wear signs and residues has allowed the author to reconstruct the functional role of this specific production, highlighting its specialized nature mostly related to the processing of animal carcasses through accurate and careful actions. The aptitude towards specialization in a tool’s function and technology shows how advanced the cognitive capacities were of the Qesem hominins. Applying functional analysis based on the determination of wear on artifacts by means of optical light microscopes, scanning electron microscopy and chemical analysis (FTIR and EDX) provides a useful and effective approach for understanding the adaptive strategies of the Qesem Cave hominins who, while facing various situations, were able to find thoughtful solutions for different needs.
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At least six different Homo species populated the World during the latest Pliocene to the Pleistocene. The extinction of all but one of them is currently shrouded in mystery, and no consistent explanation has yet been advanced, despite the enormous importance of the matter. Here, we use a recently implemented past climate emulator and an extensive fossil database spanning 2,754 archaeological records to model climatic niche evolution in Homo. We find statistically robust evidence that the three Homo species representing terminating, independent lineages, H. erectus, H. heidelbergensis, and H. neanderthalensis, lost a significant portion of their climatic niche space just before extinction, with no corresponding reduction in physical range. This reduction coincides with increased vulnerability to climate change. In the case of Neanderthals, the increased extinction risk was probably exacerbated by competition with H. sapiens. This study suggests that climate change was the primary factor in the extinction of Homo species.
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Here, we present a metric and morphological study of the molar remains from the Montmaurin-La Niche mandible by means of microcomputed tomography. According to the last analysis, based on the combination of geomorphological and paleontological data, the level bearing this human mandible probably corresponds to the marine isotope stages (MIS) 7. These data place the Montmaurin-La Niche in a chronologically intermediate position between the Neanderthals and the Middle Pleistocene fossils (e.g., Sima de los Huesos, la Caune de l’Arago). A recent study has revealed that while the mandible is more closely related to the Early and Middle Pleistocene African and Eurasian populations, the morphology of the outer enamel surfaces of its molars is typical of the Neanderthal linage. The data presented here are in line with this finding because the morphology of the enamel-dentine junction of the molars is similar to that of Neanderthals, whereas the absolute and relative enamel thickness values (2D and 3D) are closer to those exhibited by some Early Pleistocene hominins. Moreover, the pulp cavity morphology and proportions are in concordance with the Neanderthal populations. Our results strengthen the hypothesis that the settlement of Europe could be the result of several migrations, at different times, originated from a common source population. Thus, the variability in the European Middle Pleistocene populations (e.g., Montmaurin, Sima de los Huesos, Arago, Mala Balanica) could indicate different migrations at different times and/or population fragmentation, without excluding the possible hybridization between residents and new settlers.
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Quina scrapers are well-known components of the European Middle Paleolithic Mousterian. A similar production process was detected within the lithic assemblages of the Levantine Acheulo-Yabrudian (∼400–200 ka). This study combines the results of use-wear and raw material analyses of 75 Quina scrapers and 133 demi-Quina scrapers from the Acheulo-Yabrudian site Qesem Cave, Israel, aimed at interpreting the function of Quina and demi-Quina at Qesem Cave, the considerations affecting the lithic choices involved in their production, and the behavioral and evolutionary implications. Each scraper was examined for use-wear and was assigned to a flint type and potential geologic source(s). Our results demonstrate a selective pattern of exploitation of flint which does not originate from the local Turonian outcrops, specifically for the manufacture of Quina and demi-Quina scrapers. This suggests a thoughtful, well-planned effort, taking into consideration the flint traits and future function. This pattern repeats itself throughout time, reflecting a high degree of familiarity with the potential sources located around the cave, implying the existence of knowledge transmission mechanisms concerning the location of specific flint sources and their acquisition methods, as well as concerning exploitation preferences.
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This study presents the dental remains discovered at Manot Cave (MC), Western Galilee, Israel. The cave contains evidence for human occupation during the Early Upper Paleolithic period (46-33 ka) mainly of Early Ahmarian (∼46-42 ka) and Levantine Aurignacian (∼38-34 ka) cultural levels. Six teeth (three deciduous and three permanent) were found at the site, of which four could be thoroughly analyzed. The morphology of the teeth was qualitatively described and analyzed using traditional and geometric morphometric methods. A large comparative sample was used in order to assess the morphological affiliation of the Manot specimens with other Homo groups. The results provided equivocal signals: the upper first premolar (MC-9 P3) is probably modern human; the upper deciduous second molar (MC-10 dm2) and the upper second permanent molar (MC-8 M2) might be modern humans; the lower second deciduous molar (MC-7 dm2) might be Neanderthal. Owing to the small sample size and the almost total lack of distinctive characteristics, our outcome could not supply conclusive evidence to address the question of whether Manot Aurignacian population came from Europe or descended from the local Ahmarian population.
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The purposeful production of small flakes is integral to the lithic variability of many Middle Pleistocene sites. Inhabitants of the Acheulo-Yabrudian site of Qesem Cave, Israel, systematically recycled ‘old’ discarded blanks and tools, using them as cores for the production of small sharp tools with distinct technological features. These recycling end-products were produced in significant quantities throughout the human occupation of Qesem Cave, and their outstanding state of preservation made possible a functional analysis with residue detection using Fourier transform infrared spectroscopy (FTIR) and energy-dispersive X-ray spectroscopy (SEM-EDX). An experimental program accompanying the study tested the efficiency of each tool category, and a reference collection of the organic remains was assembled. Our integrated results show that small flakes were used mainly to process soft to medium animal material through precise cutting activities that required accurate longitudinal motions. Several items show clear and exclusive contact with bone while others were used for designated steps in hide treatment processes. Plant and tuber processing are also evidenced although to a lesser extent. We show that the end products of recycling ‘old’ flakes reflect preconceived technological and functional characteristics produced in a targeted manner to obtain specific tools designated for anticipated practical tasks. We demonstrate the complementary role of the products of recycling within the Qesem Cave tool- kits alongside larger tools in assisting early humans in the different stages of processing animal materials. Moreover, use-wear and residue evidence indicates that Qesem hominins differentiated their activities across space in the cave. We argue that the meticulous realization of specific tasks and the deliberate, repetitive, and skilled production of tools of different sizes and shapes is one characteristic of the new mode of adaptation practiced by Acheulo-Yabrudian hominins in the Levant in order to better manipulate the available resources following the disappearance of mega-herbivores. ©
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Earliest modern humans out of Africa Recent paleoanthropological studies have suggested that modern humans migrated from Africa as early as the beginning of the Late Pleistocene, 120,000 years ago. Hershkovitz et al. now suggest that early modern humans were already present outside of Africa more than 55,000 years earlier (see the Perspective by Stringer and Galway-Witham). During excavations of sediments at Mount Carmel, Israel, they found a fossil of a mouth part, a left hemimaxilla, with almost complete dentition. The sediments contain a series of well-defined hearths and a rich stone-based industry, as well as abundant animal remains. Analysis of the human remains, and dating of the site and the fossil itself, indicate a likely age of at least 177,000 years for the fossil—making it the oldest member of the Homo sapiens clade found outside Africa. Science , this issue p. 456 ; see also p. 389
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If we restrict the use of Homo sapiens in the fossil record to specimens which share a significant number of derived features in the skeleton with extant H. sapiens , the origin of our species would be placed in the African late middle Pleistocene, based on fossils such as Omo Kibish 1, Herto 1 and 2, and the Levantine material from Skhul and Qafzeh. However, genetic data suggest that we and our sister species Homo neanderthalensis shared a last common ancestor in the middle Pleistocene approximately 400–700 ka, which is at least 200 000 years earlier than the species origin indicated from the fossils already mentioned. Thus, it is likely that the African fossil record will document early members of the sapiens lineage showing only some of the derived features of late members of the lineage. On that basis, I argue that human fossils such as those from Jebel Irhoud, Florisbad, Eliye Springs and Omo Kibish 2 do represent early members of the species, but variation across the African later middle Pleistocene/early Middle Stone Age fossils shows that there was not a simple linear progression towards later sapiens morphology, and there was chronological overlap between different ‘archaic’ and ‘modern’ morphs. Even in the late Pleistocene within and outside Africa, we find H. sapiens specimens which are clearly outside the range of Holocene members of the species, showing the complexity of recent human evolution. The impact on species recognition of late Pleistocene gene flow between the lineages of modern humans, Neanderthals and Denisovans is also discussed, and finally, I reconsider the nature of the middle Pleistocene ancestor of these lineages, based on recent morphological and genetic data. This article is part of the themed issue ‘Major transitions in human evolution’.
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Reconstructing detailed aspects of the lives of Lower Palaeolithic hominins, who lived during the Middle Pleistocene, is challenging due to the restricted nature of the surviving evidence, predominantly animal bones and stone tools. Qesem Cave, Israel (420e200 ka) is a site that has produced evidence for a wealth of innovative features including controlled use of fire, represented by a repeatedly used hearth. Numerous charred bone and stone tools as well as wood ash have been found throughout the ten metres of archaeological deposits. Here, we describe the presence of a range of potentially inhaled, and ingested, materials extracted from samples of dental calculus from the Qesem Cave hominins. These finds offer an insight into the environment in and around the cave, while micro-charcoal highlights the need for smoke management in enclosed environments. Plant fibres and a phytolith may be evidence of oral hygiene activities or of using the teeth to work raw materials. Starch granules and chemical compounds provide a direct link to ingested plant food items. This extends the evidence for consumption of plant foods containing essential nutrients including polyunsaturated fatty acids and carbohydrates, into the Lower Palaeolithic. Together, these results represent a significant breakthrough towards a better understanding of Middle Pleistocene dietary breadth and highlight some of the challenges facing the adoption of the habitual use of fire for cooking by the Qesem Cave hominins, as well as offering an insight into their ecological knowledge and technological adaptability.
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Qesem Cave is a Middle Pleistocene site situated 12 km east of the Mediterranean coast of Tel Aviv, Israel. It is attributed to the Acheuleo-Yabrudian Cultural Complex (AYCC) of the late Lower Paleolithic period, dated to ca. 420–200 ka. This site exhibits a unique prehistoric sequence where the Amudian blade dominated industry is the main cultural component, however the scraper-dominated Yabrudian industry is also represented in distinct contexts at the cave. The chronology established by TL applied on burnt flints, ESR/U-series on herbivorous teeth and U-series on spelothems, suggests that Qesem Cave is one of the oldest sites yielding such a blade industry and a fully-fledged trajectory of Quina scrapers production.
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Dental anthropologists focus on the variation around a commonly shared pattern, a variation expressed by differences in tooth size and morphology. This book centers on the morphological characteristics of tooth crowns and roots that are either present or absent in any given individual and that vary in frequency among populations. These nonmetric dental traits are controlled largely by genetic factors and provide a direct link between extinct and extant populations. The book illustrates more than thirty tooth crown and root traits and reviews their biological and genetic underpinnings. From a database of more than 30,000 individuals, the geographic variation of twenty-two crown and root traits is graphically portrayed. A global analysis of tooth morphology shows both points of agreement and disagreement with comparable analyses of genetic and craniometric data. These findings are relevant to the hotly contested issue of timing and geographic context of modern human origins.
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The procurement and selection of raw materials for producing different stone tools in the past provide invaluable insights into hominid technological capabilities and behavior. Flint has been extensively studied to document its sources, tool production, use, and recycling. Less is known about the procurement strategies used for obtaining the raw materials. Our approach is based on the concentration of cosmogenic in situ produced 10Be within the flint. As this is depth dependent, flint material collected from the surface can be differentiated from fint collected at depths or from special environments which protected the flint from cosmic radiation. 10Be concentrations in different tool types from the Lower Paleolithic strata of Qesem cave showed that the raw materials for large scrapers and hand-axes were obtained from deep buried material or recently exposed material. The smaller blades showed a larger distribution of 10Be that resembles the concentrations of 10Be in flint nodules collected from the soil surface around the cave. This is consistent with the observation that the large scrapers and handaxes were re-sharpened. Therefore some 400,000 years ago the Qesem cave inhabitants possessed a detailed knowledge of the resources, and the capability to procure appropriate raw materials for specific tool types.
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Qesem Cave is a Middle Pleistocene site situated 12 km east of the Mediterranean coast of Tel Aviv, Israel. It is attributed to the Acheuleo-Yabrudian Cultural Complex (AYCC) of the late Lower Paleolithic period, dated to ca. 420–200 ka. This site exhibits a unique prehistoric sequence where the Amudian blade dominated industry is the main cultural component, however the scraper-dominated Yabrudian industry is also represented in distinct contexts at the cave. The chronology established by TL applied on burnt flints, ESR/U-series on herbivorous teeth and U-series on spelothems, suggests that Qesem Cave is one of the oldest sites yielding such a blade industry and a fully-fledged trajectory of Quina scrapers production.
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The lithic assemblages recovered from Qesem Cave can be divided into two distinct industries, the Amudian and the Yabrudian, based mostly on different proportions of blades and scrapers. The assemblages can also be divided chronologically and by area within the site, such as the shelf, and the hearth area. The goal of our study is to try to determine whether the assemblages vary in their use of the landscape and its resources. Were the same raw materials used in each case, or do the assemblages differ, either in proportions or in sources exploited? We have therefore undertaken an examination of a sample of approximately 6000 pieces, divided fairly equally between three assemblages: the Amudian and the Yabrudian from the shelf area, and the hearth area (which is Amudian). We begin by categorizing each distinct raw material as to rock type, and, since most of them are varieties of flint, distinguish individual types based on criteria such as colour, cortex characteristics, homogeneity, and any visible fossils. To date we have identified 51 raw material types, some of which are varieties of each other which will be grouped together later. Each lithic piece examined is recorded as to typological category, assigned a raw material type, and weighed. We are thus able to examine the differences in proportions of use of the raw material types in each assemblage, by typological category, and in terms of number of pieces and of weight of material used. This paper reports only on the data concerning numbers of pieces, and shows that some raw materials were selected for specific types of tools or technological requirements. In addition, we have started field work aimed at locating the sources of the flint varieties. We have located 15 potential sources, most within 5 km of the site, but also two located more than 15 km away. Comparison of samples from the geologic sources with samples from the site assemblages has allowed us to tentatively suggest the origins of 46 of the 51 raw material types. This gives us an idea of the extent of the territory exploited by the hominins at Qesem, how different raw materials were used for different purposes, and how use of these resources and territory differs between assemblages.
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Seventeen Middle Pleistocene crania from the Sima de los Huesos site (Atapuerca, Spain) are analyzed, including seven new specimens. This sample makes it possible to thoroughly characterize a Middle Pleistocene hominin paleodeme and to address hypotheses about the origin and evolution of the Neandertals. Using a variety of techniques, the hominin-bearing layer could be reassigned to a period around 430,000 years ago. The sample shows a consistent morphological pattern with derived Neandertal features present in the face and anterior vault, many of which are related to the masticatory apparatus. This suggests that facial modification was the first step in the evolution of the Neandertal lineage, pointing to a mosaic pattern of evolution, with different anatomical and functional modules evolving at different rates.
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Observable differences among craniomandibular specimens from Dmanisi, Republic of Georgia have resulted in controversy regarding the number of represented taxa. In this study, the hominin dental remains from Dmanisi are analyzed to help determine whether the variation present in the Georgian assemblage likely reflects interspecific or intraspecific variation, including sexual dimorphism. Metric variation is assessed with resampling methodology by comparing the Dmanisi teeth with reference samples of extant hominoids. Comparisons were also made between the variation observed in the Georgian sample and that of Paranthropus boisei. The results provide evidence for the taxonomic homogeneity of the D211/D2282 and D2735/D2700 skulls. Whether D2600 should be included within this sample or considered distinct is less clear as the distal molar crown dimensions of this robust specimen are notably larger than the other Georgian fossils. Nonetheless, it is argued that the metric size variation expressed by the Dmanisi dental sample, including D2600, is insufficient to reject the null hypothesis of a single species.
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Trigonid crest patterning in lower molars is distinctive among Late Pleistocene hominins such as Homo neanderthalensis, fossil Homo sapiens and modern humans. In this paper, we present an examination of trigonid crest patterning in the Middle Pleistocene permanent lower molar sample (n=62) of Homo heidelbergensis from Sima de los Huesos (SH). Crest expression was assessed from 3D models of the enamel and the dentine surfaces that were produced using micro-computed tomography (microCT). The aims of our analysis are to: 1) characterize the pattern of trigonid crest expression at the outer enamel and enamel-dentine junction surfaces (OES and EDJ) of the SH sample, 2) evaluate the concordance of expression between both surfaces, and 3) place trigonid crest variation in the SH sample into a phylogenetic context. Our results reveal a greater variability in the expression of trigonid crests at the EDJ (14 types) compared to the OES (4 types). Despite this variability, in almost all cases the expression of a continuous mid-trigonid or distal crest at the OES corresponds with the expression of a continuous mesial/mid-trigonid or distal trigonid crest, respectively, at the EDJ. Thus, it is possible to predict the type of trigonid crest pattern that would be at the OES in the case of partially worn teeth. Our study points to increased variability in trigonid crest expression in M3s compared to M1s and M2s. Moreover, our analysis reveals that the SH sample matches broadly the trigonid crest patterns displayed by H.neanderthalensis and differs from those exhibited by H.sapiens, particularly in the almost constant expression of a continuous middle trigonid crest at the EDJ. However, SH hominins also exhibit patterns that have not been reported in H.neanderthalensis and H.sapiens samples. Other aspects of the variability of the trigonid crest expression at the dentine are presented and discussed.
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Thesis (Ph. D.)--Arizona State University, 2002. Includes bibliographical references (p. [210]-238). Photocopy.
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Morphometric relationships among late middle and early late Pleistocene fossil hominids from southwestern Asia are examined utilizing multivariate analyses of frontal bones. Particular emphasis is placed on assessing the interaction of size and shape factors in determining relationships among these specimens. Results of these analyses can be summarized in five points. First, techniques which maximize pattern recognition based primarily on shape factors provide the most reliable information pertinent to phylogenetic relationships among these hominids. Second, the Zuttiyeh specimen does not exhibit a greater similarity to the early “modern” Skhu¯l/Qafzeh hominids than to Levantine Neandertals. Third, the Shanidar Neandertals do not cluster closely with Levantine archaic humans. Fourth, the Mousterian-associated Skhu¯l/Qafzeh hominids exhibit distinct similarities to archaic humans, but the Skhu¯l hominids give no unequivocal indication of being “hybrids” between the Qafzeh people and Neandertals. Finally, clear patterns of change occur in frontal bone morphology from the early “modern” (Mousterian-associated) Skhu¯l/Qafzeh hominids to early Upper Paleolithic-associated humans from the Levant.
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Le present article concerne l'importance des sites et des fossiles decouverts au Proche-Orient ou deux groupes d'Homo sapiens, Neandertaliens et Hommes modernes, ont ete identifies dans les niveaux mousteriens du Paleolithique moyen. Les principaux problemes de chronostratigraphie et leurs interpretations sont, ici, discutes. Les donnees confirment la presence des lignees des Neandertaliens du Proche-Orient et des Hommes modernes du Levant au moins depuis 150 Ky., sur la base de Tabun 1 et de Zuttiyeh, consideres, respectivement, comme les plus anciens representants de chacune d 'elles. La coexistence des deux groupes, ayant eu le meme milieu culturel pendant tout le Paleolithique moyen, est demontree ; la possibilite de metissage ne peut etre exclue. La presence effective des Hommes modernes dans la region consideree est comparee avec l'information disponible pour l'Asie, l'Europe et l'Afrique, en insistant particulierement sur l'origine problematique des Hommes modernes et leur dispersion.
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The origin of anatomically modern Homo sapiens and the fate of Neanderthals have been fundamental questions in human evolutionary studies for over a century. A key barrier to the resolution of these questions has been the lack of substantial and accurately dated African hominid fossils from between 100,000 and 300,000 years ago. Here we describe fossilized hominid crania from Herto, Middle Awash, Ethiopia, that fill this gap and provide crucial evidence on the location, timing and contextual circumstances of the emergence of Homo sapiens. Radioisotopically dated to between 160,000 and 154,000 years ago, these new fossils predate classic Neanderthals and lack their derived features. The Herto hominids are morphologically and chronologically intermediate between archaic African fossils and later anatomically modern Late Pleistocene humans. They therefore represent the probable immediate ancestors of anatomically modern humans. Their anatomy and antiquity constitute strong evidence of modern-human emergence in Africa.
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We present here the results of a U–Th dating project at Qesem Cave, a Middle Pleistocene, late Lower Paleolithic site in Israel. It provides 54 new MC-ICP-MS U–Th ages for speleothems from the cave. The results indicate that human occupation started sometime between ∼420 and 320 ka and ended between 220 and 194 ka. A survey of dates from culturally similar sites in the Levant indicates that the general range of ca. 400–ca. 200 ka is an appropriate estimate for the life span of the Acheulo-Yabrudian Cultural Complex (AYCC).
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Mesiodistal and buccolingual crown dimensions of the right deciduous teeth of 133 white children were analyzed for information on sexual dimorphism and sex discrimination using discriminant analysis. Even though consistent differences were found for only 15 out of 20 paired measurements, five of them significant at p = 0.05 or better, discriminant analysis showed the possibility of correctly sexing up to 75% of the juvenile sample, using a maximum of seven deciduous teeth.
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Comparative analyses between Gran Dolina-TD6 and Sima de los Huesos (SH) dental samples from Sierra de Atapuerca (Spain) suggest that hominins represented by these samples belonged to two very different populations and even to distinct paleospecies. Gran Dolina-TD6 hominins (Homo antecessor) have been dated to about 0.8 million years ago (mya), whereas a new radiometric dating of the SH hominins (Homo heidelbergensis) suggests an interval of 0.4–0.5 mya for these hominins. Current results as well as the relative temporal closeness between the populations represented by Gran Dolina-TD6 and SH favour a replacement scenario hypothesis (or possibly a crossbreeding scenario) for the Europeans during the early Middle Pleistocene. Although the information available from the European Lower Pleistocene populations is limited (Ceprano and Gran Dolina-TD6), current data are noteworthy and their combination with archaeological evidence will stimulate future discussions on the dynamics of the first European settlements.
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The Zuttiyeh hominin craniofacial fossil was discovered in Israel in 1925. Radiometric dates and the archaeological context (Acheulo-Yabrudian) bracket the associated cave layers to between 200 and 500 ka (thousands of years ago), making it one of the earliest cranial fossils discovered in the Near East thus far. Its geographic position, at the corridor between Africa and Eurasia, in combination with its probable Middle Pleistocene date make it a crucial specimen for interpreting later human evolution. Since its discovery, qualitative descriptive and traditional morphometric methods have variously suggested affinities to Homo erectus (Zhoukoudian), Homo neanderthalensis (Tabun), and early Homo sapiens (Skhul and Qafzeh). To better determine the taxonomic affinities of the Zuttiyeh fossil, this study uses 3D semilandmark geometric morphometric techniques and multivariate statistical analyses to quantify the frontal and zygomatic region and compare it with other Middle to Late Pleistocene African and Eurasian hominins. Our results show that the frontal and zygomatic morphology of Zuttiyeh is most similar to Shanidar 5, a Near East Neanderthal, Arago 21, a European Middle Pleistocene hominin, and Skhul 5, an early H. sapiens. The shape differences between archaic hominins (i.e., Homo heidelbergensis and H. neanderthalensis) in this anatomical region are very subtle. We conclude that Zuttiyeh exhibits a generalized frontal and zygomatic morphology, possibly indicative of the population that gave rise to modern humans and Neanderthals. However, given that it most likely postdates the split between these two lineages, Zuttiyeh might also be an early representative of the Neanderthal lineage. Neanderthals largely retained this generalized overall morphology, whereas recent modern humans depart from this presumably ancestral morphology.
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The systematic excavation of the Sima de los Huesos (SH) site in Sierra de Atapuerca (Burgos, Spain) has yielded the largest hominin collection worldwide for the Middle Pleistocene. The dental sample now consists of more than 500 teeth that provide exceptional opportunities to define the dental morphological pattern of a Middle Pleistocene population as well as develop hypotheses about the origins of the Neanderthals. The dental collection has now increased to over 533 specimens (525 permanent and 8 deciduous teeth), necessitating new morphological assessments. Thus, we present a detailed morphological description of the SH permanent dentition recovered up to 2007, accomplishing comparisons with European Middle Pleistocene hominins, Neanderthals, and early and contemporary Homo sapiens. We find that SH dentitions present all the morphological traits that, either in their degree of expression, frequency, or particular combination, are usually considered as typical of Homo neanderthalensis. This study ratifies the deep roots of the Neanderthal lineage in the Middle Pleistocene of Europe. In addition, SH teeth are morphologically "more Neanderthal" than other penecontemporaneous Middle Pleistocene samples such as Mauer or Arago, and even more derived than some classic Neanderthal samples. Thus, our study would not sustain the linearity of the accretion process hypothesized for the origins of the Neanderthals, and we suggest that other evolutionary models and scenarios should be explored for the Middle and Upper Pleistocene of Europe. We propose that more than one hominin lineage may have coexisted during the Middle Pleistocene in Europe.
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This study presents a description and comparative analysis of Middle Pleistocene permanent and deciduous teeth from the site of Qesem Cave (Israel). All of the human fossils are assigned to the Acheulo-Yabrudian Cultural Complex (AYCC) of the late Lower Paleolithic. The Middle Pleistocene age of the Qesem teeth (400-200 ka) places them chronologically earlier than the bulk of fossil hominin specimens previously known from southwest Asia. Three permanent mandibular teeth (C(1) -P(4) ) were found in close proximity in the lower part of the stratigraphic sequence. The small metric dimensions of the crowns indicate a considerable degree of dental reduction although the roots are long and robust. In contrast, three isolated permanent maxillary teeth (I(2) , C(1) , and M(3) ) and two isolated deciduous teeth that were found within the upper part of the sequence are much larger and show some plesiomorphous traits similar to those of the Skhul/Qafzeh specimens. Although none of the Qesem teeth shows a suite of Neanderthal characters, a few traits may suggest some affinities with members of the Neanderthal evolutionary lineage. However, the balance of the evidence suggests a closer similarity with the Skhul/Qafzeh dental material, although many of these resemblances likely represent plesiomorphous features.
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The nearly ubiquitous presence of a continuous crest connecting the protoconid and metaconid of the lower molars (often referred to as the middle trigonid crest), is one of several dental traits that distinguish Homo neanderthalensis from Homo sapiens. This study examined variation in trigonid crest patterns on the enamel and dentine surfaces to (1) evaluate the concordance between the morphology of trigonid crests at the inner dentine and the outer enamel surfaces; (2) examine their developmental origin(s); and (3) examine trait polarity through comparison with Australopithecus africanus and Pan. The sample included 73 H. neanderthalensis, 67 contemporary H. sapiens, 5 A. africanus, and 24 Pan lower molars. Results indicate general agreement in the morphology observed on the dentine and enamel surfaces. All but one H. neanderthalensis molar shows some trigonid crest development, whereas trigonid crests occur in low frequency in contemporary humans. Pan and A. africanus both also show high frequencies of a continuous trigonid crest. However, the origin of the trigonid crest differs among groups. H. neanderthalensis uniquely possesses a 'middle' trigonid crest that originates from the mesial accessory ridge of one or both cusps. Based on our results we suggest that presence of a continuous middle trigonid crest at the dentine surface is primitive and the lack of any trigonid crest is derived. Genetic drift may explain the high frequency of trigonid crests in H.neanderthalensis. However, H. neanderthalensis still appears to be derived relative to Pan and A. africanus in its high frequency of the mesial-mesial trigonid crestconfiguration.
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Most research on Neandertal teeth has focused on shovel shaped incisors and/or taurodont molars. In the past 15 years there has been a renewed interested in Neandertal dental morphology, especially with regard to how they compare to recent and fossil modern humans. However, no complete description of Neandertal dental morphology has been published since the mid-1950s. Many more Neandertals and other fossil hominins have been discovered since then and are available for a comparative study. This paper provides a description of Neandertal dental morphology and places that morphology in a comparative fossil hominin context. It differs from previous work by focusing on fossil hominin variation (as opposed to contemporary modern human variation) and provides a comparative baseline in which Neandertal dental morphology can be assessed. The four comparative samples include European and West Asian Neandertals, European non-Neandertal archaics, South African/West Asian early modern humans and European early modern humans (e.g., Upper Paleolithic). A mean measure of divergence analysis shows that Neandertals are significantly different from early modern human groups, being four times more divergent from Afro-Asian and European early modern human samples than the early modern human samples are from each other. Moreover, Neandertals are more divergent from early modern Europeans than they are from the early modern Afro-Asians. Contrary to the results of a previous study they are significantly divergent from non-Neandertal archaics. The implications for these results are discussed.
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Among primitive peoples dental attrition appears to be a natural phenomenon. Often the degrees and kinds of tooth wear vary from population to population. This variability is possibly related to certain material aspects of culture such as diet, food preparation techniques and tool usage. In order to learn more about these relationships, extensive cross cultural comparisons must be made. This paper reports on a study of dental attrition among skeletal remains of North American Indians from three areas: California, the Southwest and the Valley of Mexico. A method of comparing worn teeth of these populations was devised so several characteristics of the teeth and supporting bone could be examined by population. This study showed significant differences in type and degree of wear among the three groups as well as differences between sexes within each population. A positive correlation between tooth wear and cultural factors was found. Dietary specialization and division of labor appear to be responsible for the degree and type of wear found in this sample. Further studies of this type are planned to expand the sample size and, if the new data support these correlations, valuable information about human–environmental relationships can be gained.